Received: 16 September 2017 | Revised: 20 June 2018
DOI: 10.1111/jbi.13422
RESEARCH PAPER
Islands in the trees: A biogeographic exploration of epiphyte‐
dwelling spiders
Francisco Emmanuel Méndez-Castro1,2
Dinesh Rao2
1
Ecological Plant Geography, Faculty of
Geography, University of Marburg,
Marburg, Germany
2
Instituto de Biotecnología y Ecología
Aplicada (INBIOTECA), Universidad
Veracruzana, Xalapa, México
Correspondence
Francisco Emmanuel Méndez-Castro,
Deutschhausstraße 10, 35032 Marburg,
Germany.
Email: femendez@icloud.com
Funding information
Universidad Veracruzana, Grant/Award
Number: PROMEP/103.5/10/7748;
CONACYT, Grant/Award Number: 307402/
251146
Editor: Triantis Kostas
Journal of Biogeography. 2018;1–10.
| Accepted: 30 June 2018
| Maaike Y. Bader1 | Glenda Mendieta-Leiva1 |
Abstract
Aim: Epiphytic plants are isolated from each other by nonhabitat canopy elements
and are thus expected to act as islands, the biodiversity of their inhabitants (e.g.,
spiders) conforming to island biogeographic predictions of species‐richness patterns.
Although it has been shown that arthropod diversity decreases with decreasing epi-
phyte size, the effects of isolation have not yet been addressed. We studied the
joint effect of isolation, spatial position, and size of epiphytic plants (canopy islands)
on species richness, relative rareness, and similarity of spider communities.
Location: A shade‐coffee plantation in central Veracruz, Mexico.
Taxon: Spiders (Araneae), vascular epiphytes (Bromeliaceae, Piperaceae, Orchi-
daceae, Araceae, Pteridophyta).
Methods: We collected all canopy islands occurring on three trees and recorded
their three‐dimensional spatial position. In the laboratory, we disassembled the
plants and collected all spiders present. We analysed the effects of island size, isola-
tion, and spatial position on the species richness and the relative rareness of spider
communities using generalized linear models (GLM), on the distribution of different
spider guilds using a CCA, and on community composition using a permutational
multivariate analysis of variance (Permanova). Isolation and spatial position were
addressed using five distance measures representing isolation from different poten-
tial species sources. We tested how the similarity in spider community composition
changed with the distance between islands using Mantel correlograms.
Results: As predicted by island biogeography theory, spider species richness
increased with canopy‐island size and decreased with isolation. In comparison to
island size, the effect of isolation was weak, though significant. Relative rareness
was hardly affected by island size but more by isolation. Compositional similarity
was affected by island size and decreased with increasing spatial distance up to ca.
4 m. Guilds separated along the main CCA axes, this ordination being driven by epi-
phyte size and position.
Main conclusions: Epiphytic plants behaved like canopy islands in that their size
and isolation influenced the diversity and composition of spider communities. How-
ever, the effect of isolation was only a fraction of that of island size, perhaps
because spatial relationships are taxon‐specific. This may be due to differences in
hunting behaviour and dispersal capacities, for example, between guilds of hunting
wileyonlinelibrary.com/journal/jbi © 2018 John Wiley & Sons Ltd | 1
2 |
and web‐building spiders, which is a dimension deserving more attention. For a bet-
ter understanding of biogeographic principles driving the diversity of canopy island
inhabitants, further research on this topic should include position and isolation, at
scales matching the mobility of different functional groups, as part of their explana-
tory variables.
KEYWORDS
epiphytes, habitat islands, Island biogeography, spatial patterns, species richness, spiders
1 | INTRODUCTION
According to the equilibrium theory of island biogeography (ETIB),
species numbers on islands are primarily determined by the interac-
tion of island size and isolation (Cox, Moore, & Ladle, 2016;
MacArthur & Wilson, 1967; Whittaker & Fernández‐Palacios, 2008).
Larger and less isolated islands are expected to harbour more spe-
cies than smaller and more isolated ones. Islands do not need to be
surrounded by water; any habitat patch embedded in a nonhabitat
matrix can be considered an island and species numbers in such
habitat islands should conform to the size–isolation pattern predicted
by the ETIB (Adams, Schnitzer, & Yanoviak, 2016; Öckinger, Franzén,
Rundlöf, & Smith, 2009; Prugh, Hodges, Sinclair, & Brashares, 2008).
Epiphytic plants in the canopy of tropical forests can be considered
as islands since they are discrete habitats separated from each other
by a dissimilar matrix. These canopy plants form habitat islands for
many arthropods and other organisms (Richardson, 1999) and are
arranged in “archipelagos” in a three‐dimensional and discontinuous
habitat space (Bader, van Dunné, & Stuiver, 2000).
In accordance with the ETIB, larger epiphytes have been found
to host a greater number of arthropod species such as ants, beetles,
springtails, and spiders (Méndez‐Castro & Rao, 2014; Rodgers &
Kitching, 2011; Stuntz, Ziegler, Simon, & Zotz, 2002; Wardle et al.,
2003). However, the effect of isolation on faunal diversity is difficult
to demonstrate (Barton, Evans, Foster, Cunningham, & Manning,
2017; Jiménez‐Valverde, Baselga, Melic, & Txasko, 2009; Wu, Si,
Didham, Ge, & Ding, 2017). So far, isolation effects (interisland dis-
tance) on canopy fauna have been studied only once, for canopy soil
communities in real and artificial soil accumulations in New Zealand,
where no significant effects were detected (Wardle et al., 2003).
For most islands, isolation is defined as the linear distance(s) of a
given island to one or several species sources (mainland or neighbour
islands). Unlike distances between real islands and terrestrial habitat
fragments, distances between canopy islands are defined in a three‐
dimensional space (Bader et al., 2000). It is unclear whether isolation
within this space plays a role in determining species distributions.
One reason for this uncertainty is the multitude of potential species
sources, such as neighbouring epiphytes, the understorey or the sur-
rounding trees. The effect of epiphyte spatial distribution has been
addressed previously, but only considering vertical stratification,
which is generally found to be important for the composition of
canopy fauna (Aikens & Buddle, 2012; Gonçalves‐Souza, Almeida‐
MÉNDEZ‐CASTRO ET AL.
Nieto, & Romero, 2011; Méndez‐Castro & Rao, 2014; Pinzon,
Spence, & Langor, 2013).
In this study, we analysed the combined effect of epiphyte size,
spatial position and isolation on the species richness, the relative pro-
portion of rare species, and the community similarity of epiphyte‐inha-
biting spiders. Spiders and epiphytes have a mutualistic relationship in
which the plants provide spiders with a suitable habitat for foraging,
reproduction, egg laying, and finding shelter (Gonçalves‐Souza,
Brescovit, Rossa‐Feres, & Romero, 2010; Scheffers, Edwards, Dies-
mos, Williams, & Evans, 2014), and spiders reduce herbivore popula-
tions through predation and can make nutrients available to the plants
(e.g., via excretion) (Gonçalves, Mercier, Mazzafera, & Romero, 2011;
Romero, Mazzafera, Vasconcellos‐Neto, & Trivelin, 2006). Spiders are
highly sensitive to subtle changes in habitat structure (Gonçalves‐
Souza et al., 2011) and microclimatic conditions and they show active
habitat selection (Omena & Romero, 2008; Rao, 2017). Because of this
sensitivity, habitat diversity is expected to positively affect spider
species numbers even at small spatial scales (Aikens & Buddle, 2012;
Barton et al., 2017; De Mas, Chust, Pretus, & Ribera, 2009). On top of
being selective and sensitive towards the physical structure of their
habitat, spiders display a variety of foraging strategies and can be
classified into functional guilds (Cardoso, Pekár, Jocqué, & Coddington,
2011), which differ in their mobility (e.g., web builders vs. active
hunters). These differences are expected to work as behavioural filters
modulating the effects of island size and isolation on spider diversity
at the community level (Cobbold & MacMahon, 2012).
Although spiders can move between epiphytes along branches
or, depending on the species, through the air (Mestre, Bucher, &
Entling, 2014; Wolff, Schneider, & Gorb, 2014), the space between
the epiphytes contains only a fraction of the spider occurrences
observed in epiphyte‐rich trees (Angelini & Silliman, 2014; Cruz‐
Angón, Baena, & Russell, 2009) and is therefore considered as a
nonhabitat matrix in this study.
We aimed to test to what extent the predictions of the ETIB with
respect to size and isolation effects on biodiversity are valid for spider
communities at the spatial scale of canopy ecosystems. We hypothe-
sized that: (a) epiphyte size will have a positive effect on spider species
numbers, (b) isolation will have a negative effect on spider species
numbers and community similarity, (c) more isolated epiphytes will
contain a higher proportion of species that are rare in the overall sam-
pling, and (d) spiders will show specific responses towards island fea-
tures depending on their hunting strategies (guilds).
MÉNDEZ‐CASTRO ET AL. | 3

2 | MATERIALS AND METHODS 2.2 | Analysis

2.1 | Data collection We evaluated what proportion of the locally available spider species
pool was represented in the three trees by means of the interpola-
The study area is located in southeast Xalapa, Veracruz, Mexico. The tion and extrapolation analysis described by Chao et al. (2014),
site is a 40‐year‐old shade‐coffee plantation called “La Orduña” which assesses sample completeness based on sample coverage. We
(19°28′02″N, 96°55′45″W), a well‐known site for the study of spe- used the function iNEXT from the R package ‘iNEXT’ (Hsieh, Ma, &
cies diversity (Cruz‐Angón et al., 2009; Manson, Hernández‐Ortiz, Chao, 2016).
Gallina, & Mehltreter, 2008). We selected three trees of the domi- The full epiphyte and spider data sets included four dependent
nant species Inga jinicuil G. Don (Mimosaceae), which harbours the and 10 independent variables, out of which a subset was used for
greatest diversity of epiphytes (more than 40 species; Cruz‐Angón et analysis (data summarized in Tables 1 and 2, variable descriptions
al., 2009). Epiphytes and spiders were collected as follows: for the subset below and for the rest in Appendix S1). We tested
the collinearity of the variables using pairwise nonparametric
1. We climbed the trees using the single rope technique (Perry, 1978). Spearman correlations (corr.test function of the R package ‘psyc’;
2. We recorded three-dimensional epiphyte positions: “x” and “y” Revelle, 2017). When variables strongly correlated (r > |0.5|), we
coordinates were measured along measuring tapes, with the retained only those we considered most informative and discarded
trunk of the tree as the origin (Figure S1 in Appendix S1), and those that were redundant (Table S2 in Appendix S1). From the
the position on the “z” axis (corresponding to the distance from full set of four dependent variables (abundance, species richness,
the ground, as the trees were located on flat terrain) was mea- functional diversity, and relative rareness), we retained only spe-
sured by dropping down a measuring tape. cies richness, as the most basic diversity measure, and relative
3. We carefully collected canopy islands by removing the roots of rareness, which was least correlated (r ≤ |0.30|). In the case of
the epiphytes from the tree, after which each plant was wrapped independent variables, all island size and diversity variables (epi-
in a plastic bag to avoid the loss of mobile spiders (Stuntz et al., phyte abundance, epiphyte dry weight (log island size), epiphyte
2002; Yanoviak, Nadkarni, & Gering, 2003). groups, and habitat diversity index, Table S2 in Appendix S1)
4. We disassembled canopy islands in the laboratory and captured showed strong correlations with r > |0.6|, so that we retained only
all spiders. We collected spiders by hand and preserved them in island size. The spatial variables (branch diameter, distance from
80% ethanol for further examination. We sorted collected spiders the ground, horizontal distance, average isolation, nearest neigh-
into morphotypes according to their morphological traits and, bour distance, and nearest neighbour index, Table S2 in
when possible, we identified adults to species following taxo- Appendix S1) correlated only weakly, with r < |0.30|, except
nomic keys available in the literature (Jocqué & Dippenaar- branch diameter with distance from ground (r = −0.51) and hori-
Schoeman, 2006; Ubick, Paquin, Cushing, & Roth, 2005; World zontal distance (r = −0.69). Therefore, all spatial variables except
Spider Catalog, 2018). Spider vouchers are located at the Labora- branch diameter were maintained.
torio de Aracnología de la Facultad de Ciencias, Universidad
Nacional Autónoma de México (UNAM).
5. We dried the epiphytes for two days at 70°C and weighed them 2.2.1 | Dependent variables used
(Affeld, Sullivan, Worner, & Didham, 2008). Spider species richness corresponds to the number of species per
canopy island. The relative proportion of rare spiders per island was
Before collecting epiphytes (or “canopy islands”), we conducted a determined using the “Index of Relative Rarity” (hereafter referred
pilot analysis (nearest‐neighbour clustering) on 17 Inga jinicuil trees to
identify those with the highest structural similarity. Variables consid-
T A B L E 1 Summary of tree, canopy island (epiphyte), and spider
ered were diameter at breast height, number of branches, epiphyte
community characteristics for three trees in a shade‐coffee
load, and canopy cover. Trees analysed were at least 50 m apart from plantation in Mexico. Canopy islands could occur either as epiphyte
each other to ensure independence. Based on our previous experience mats composed of several plants or as single plants
collecting epiphyte‐dwelling spiders, spiders can last in the collection Tree Tree 1 Tree2 Tree3 Total
bags up to five days without getting damaged or suffocating (Méndez‐
Stem DBH (cm) 52 51 59
Castro & Rao, 2014). Therefore, we collected a maximum of 50 epi-
Tree height (cm) 1480 1480 1260
phytes per day to have enough time to process the samples and to
Maximum tree‐canopy radius (cm) 432 623 780
avoid accumulation of unprocessed samples. Because of the high num-
Epiphyte individuals 363 314 514 1191
ber of epiphytes per tree (Table 1), it took us about seven weeks to
Canopy islands 120 83 96 299
collect ca. 95% of the epiphytes occurring on a single tree. The remain-
Total spider species richness 94 89 99 164
ing 5% of plants corresponded to epiphytes found on the thin
branches of the outer canopy, which were not collected for safety rea- Total number of rare species 64 56 58 127

sons. We collected epiphytes from October 2013 to March 2014. Total number of spiders 2631 2736 2113 7480
4 | MÉNDEZ‐CASTRO ET AL.

T A B L E 2 Summary of canopy island (epiphyte) and spider
community characteristics for the canopy islands on three trees in a
shade‐coffee plantation in Mexico. To determine the epiphyte group
diversity, we defined six groups of epiphytes according to their
taxonomy: Bromeliaceae, Orchidaceae, Pteridophyta, Piperaceae,
Araceae, and miscellaneous (some epiphytic cacti and hemiepiphytic
Ficus). Habitat diversity is an index that combines island size and the
number of epiphyte groups
Canopy island Min Mean Max
any of them were collected. We tested whether this might have
masked isolation effects (due to communities reaching new equilibria
in between sampling campaigns) using a timed average isolation (tai),
calculated by excluding those islands that were removed by previous
sampling from the calculation of average isolation. If there would be
a significant effect of sampling time, we would expect tai to have a
higher explanatory power than ai.
We ran generalized linear models (GLM) to analyse the effect of

Number of epiphytes 1 5 40 canopy‐island size and spatial variables on spider species richness

Epiphyte group diversity (egd) 1 2 6

Spider species richness 1 8
Number of spiders 1 24 487
Branch diameter (cm) 2 14
Canopy‐island size (cis, in g) 1 170 4222
Habitat diversity (log [(cis +1) * egd]) 0.3 3.4
Nearest neighbour distance (cm) 10 70 233
Average isolation (cm) 369 524 990
as “relative rareness”), which was calculated by assigning weights to
spider species based on their abundance in the total sample and
then summing these weights per island. These summed weights are
then divided by the species richness of each canopy island so that
the proportion depends on the number of rare species but not on
species richness (Leroy, Petillon, Gallon, Canard, & Ysnel, 2012).
2.2.2 | Independent variables
Epiphyte biomass (dry weight) was used as the measure of island
size. Euclidean distances between the epiphytes were calculated
using the function vegdist of the R package ‘vegan’ (Oksanen et
al., 2017). The spatial multidimensionality of the canopy was
addressed by using five distance variables (Figure 1): (a) distance
from the ground (dg) corresponds to the distance on the “z” axis
(height) and represents isolation from the understorey; (b) horizon-
tal position (hod) represents isolation from the centre of the tree
(corresponding to the zone with the largest epiphyte biomass;
Freiberg & Freiberg, 2000; Díaz, Sieving, Peña‐Foxon, Larraín, &
Armesto, 2010; Nakanishi, Sungpalee, Sringernyuang, & Kanzaki,
2016) (c) nearest neighbour distance (nnd) is the Euclidean dis-
tance from a given canopy island to its closest neighbour and rep-
resents isolation from the nearest potential species source; (d)
nearest neighbour index (nni) represents edge to the edge (rather
than centre to centre) Euclidean distance between canopy islands
and was calculated as nni ¼ pnnd
ffiffiffiffiffiffiffiffi, where nns = nearest‐neighbour
nns
size and nnd = nearest‐neighbour distance (adaptation of an idea
proposed by Gustafson & Parker, 1994); and (e) average isolation
(ai) is the average distance of a given island to all other islands in
the same tree and summarizes the spatial position of canopy
islands in the tree.
Isolation variables were calculated considering the position of
the canopy islands in their initial configuration on the trees, before
and relative rareness. The initial models included island size, distance
from the ground, horizontal distance, average isolation, nearest
37
neighbour distance, and nearest neighbour index and all two‐way
interactions. Tree identity was included as a covariable to account
59
for intertree variation. Additionally, for tree1, we calculated these
models with ai replaced by tai. To find the minimal set of indepen-
18.1
dent variables explaining the highest amount of variation, we used
stepwise backwards model selection (drop1 function) to remove the
least significant variables. The percentage of explained variance (D2)
was calculated by dividing the residual deviance of each term (pro-
vided by the function anova) by the total deviance of the model
(Kindt & Coe, 2005).
To assess how the similarity (Bray–Curtis) between spider com-
munities correlated with the Euclidean distance between canopy
islands, we computed Mantel correlograms for each tree. The dis-
tance classes were set at 100‐cm intervals and we used only dis-
tance classes that included comparisons between all canopy islands
(distances above 700 cm were omitted). We used Spearman correla-
tions and 999 permutations. Using an analysis of similarity (function
anosim), we compared dissimilarities in spider species composition
between islands among and within trees, and with a permutational
multivariate analysis of variance (Permanova, function adonis2), we
analysed which of the tree and canopy island features explained
these dissimilarities. Both anosim and permanova use dissimilarity as
the default distance measure. We explored the specific responses of
spider guilds to island features using a canonical correspondence
analysis (CCA) based on the species per guild, plotting the guilds,
and independent variables. Guilds were assigned to spider species
according to their family (Table S2.2 in Appendix S2) following the
guild classification proposed by Cardoso et al., (2011). Anosim,
Permanova, correlograms, and CCA were all calculated using the
R package ‘vegan.’
All analyses were performed in the statistical software R version
3.3.2 (R Development Core Team, 2015).
3 | RESULTS
We collected 1,191 epiphytic plants clustered in 299 canopy islands
on three trees (Table 1). We extracted 7,480 spiders and sorted
them into 164 morphospecies representing 48 genera and 26 fami-
lies (Table S2.1 in Appendix S2). Trees harboured about 98% of the
species likely to be present in the area while each tree harboured
over 90% (Figure S3 in Appendix S1).
MÉNDEZ‐CASTRO ET AL. | 5

Axis origin

1200 Horizontal distance (hod)

3

1000 Nearest neighbor distance (nnd)

5
800

Distance to ground (dg)

Z

600
6
4

400 35 30 25 20 15 10 5

2 Spider species richness

7
200
Branch diameter
1 Spider species richness
600
400
400 200
200 0
0 -200
-200 -400 X
Y
-400 -600
-600

F I G U R E 1 Three‐dimensional scatter plot of the spatial position of canopy islands (composed of epiphytes) in a tree (tree 3) in a Mexican
coffee plantation, showing the meaning of different distance and isolation metrics. Canopy islands are represented by bubbles, bubble size
(legend 1) represents island size, and bubble colour (legend 2) refers to spider species richness. Horizontal distances to the trunk (arrow 3)
were calculated taking the axis origin as a reference point. Average isolation (inset 4) is the mean Euclidean distance from a given canopy
island to all other islands occurring on the same tree. In most cases, the nearest neighbour (arrows 5) of given canopy islands was located on
the same branch, but it could also be on a neighbouring branch. Distance from the ground (arrow 6) and branch diameter (bars 7) were
measured at the same point (the middle of the epiphyte mats). The scale of the axes is in cm; canopy island positions were recorded with this
precision

The spider families with highest abundances were the Dictynidae
(n = 1042), Pholcidae (n = 899), and Anyphaenidae (n = 897), while
the most species‐rich families were Salticidae (23 spp.), Theridiidae
(14 spp.), and Anyphaenidae (12 spp.). Around 75% of the species
were classified as rare in the total sample.
Generalized Linear Models (Table 3) showed that although island
size, distance from the ground, and average isolation were significant
drivers of spider species richness and relative rareness, their contri-
bution to the amount of explained variation differed according to
the dependent variable. Overall, the models could explain 77% of
the variation in species richness and just 13% of that in relative rare-
ness. Island size explained about 68% of the variation in species rich-
ness (Table 3, M1) but only 3.1% of the variation in relative rareness
(Table 3, M2). Only in the model for relative rareness, island size
interacted with average isolation (Figure S4C in Appendix S1). Dis-
tance from the ground was the most important explanatory variable
for relative rareness (explaining 6.2% of the variation), while for spe-
cies richness, it explained <1%. The third significant explanatory vari-
able for relative rareness was average isolation, while for species
richness, this variable was only significant in interaction with hori-
zontal distance (Figure S4B in Appendix S1), explaining only 1.7% of
the variation. Distance from the ground and horizontal distance (Fig-
ure S4A in Appendix S1) also significantly interacted in the species
richness model, but again with little explanatory power (0.6% of vari-
ation explained). Models using timed average isolation (Table S6 in
Appendix S1) retained the same set of explanatory variables as mod-
els using average isolation (Table 3). While timed isolation produced
no improvement on the explanatory power of species richness
model, it doubled the amount of explained deviance for the model
of relative rareness. However, timed isolation itself did not have a
significant effect but only modified the effect of distance from the
ground through an interaction. Therefore, we concluded that using
6 | MÉNDEZ‐CASTRO ET AL.

T A B L E 3 Generalized linear models of spider diversity on epiphytic plants in a Mexican shade‐coffee plantation. The two models differ in
their dependent variables (M1: species richness, M2: relative rareness). Shown are model estimates, p‐values, and the percentage of explained
variation (%D2) for each variable. We used the quasipoisson family for both models. Even though all combinations of island and spatial
variables and their two‐way interactions were tested, the minimal models presented here contain only the significant variables selected using
stepwise model selection
M1 M2

Species richness Relative rareness

z value Pr(z) %D 2
z value Pr(z) %D2
(Intercept) 54.7 *** – (Intercept) −32.8 *** –
Log island size (lis) 29.2 *** 68.5 Log island size (lis) 1.8 . 1.0
Distance from the ground (dg) −3.8 *** 0.8 Distance from the ground (dg) −3.9 *** 6.2
Horizontal distance (hod) −1.1 – 2.1 Average isolation (ai) 2.5 * 2.9
Average isolation (ai) 1 – 0.0 lis : ai −3.3 ** 3.1
Tree2 −7.1 *** 3.4 Total %D2 13.2
Tree3 −6.6 ***
dg : hod −3.9 *** 0.6
hod : ai −4.9 *** 1.7
Total %D2 77.1

Interactions are denoted by “:” between pairs of variables. Significance codes: <0.001 “***” 0.001 “**” 0.01 “*” 0.05 “–” >0.05.

timed isolation, that is, taking into account the changes in spatial 0.1
configuration caused by the sampling, did not help to understand
the ecological patterns in spider distributions. 0.05
Mantel correlation

The Mantel correlograms (Figure 2) showed that increasing dis-

0
tances between canopy islands decreased species similarity
(Bray Curtis) of spider communities (negative correlation) up to
-0.05
about 4 m. At larger distances, correlations turn positive and increas-
Tree 1
ing distances increase similarity. Two of the three trees conformed -0.1
Tree 2
to this pattern, but one showed a different result, with just one sig-
nificant negative correlation for distances around 1 m (Figure 2, -0.15
Tree 3
50 100 200 300 400 500 600 700
tree2). The analysis of similarity (anosim) revealed that within trees,
the mean dissimilarity (Bray–Curtis) between canopy islands was Distance class index
around 18%, which was significantly lower than the mean among
F I G U R E 2 Mantel correlograms based on the Bray–Curtis
trees 25% (Figure S5 in Appendix S1). According to the permuta- similarity of spider diversity and the Euclidean distances (in cm)
tional multivariate analysis of variance (Permanova), the dissimilarity between canopy islands (epiphytes) in three trees in a Mexican shade‐
in spider species composition between canopy islands was explained coffee plantation. Grey symbols represent nonsignificant correlations
by island size and tree identity (Figure S5 in Appendix S1). (p > 0.05)
Canonical correspondence analysis (Figure 3) organized guilds
into four groups suggesting specific responses towards island fea-
tures. Together, island size, distance from the ground, and horizontal 4 | DISCUSSION
distance explained 26% of the ordination of the guilds, with island
size as the dominant feature (variation explained, R2 = 0.22). The Trees can be islands for small cursorial animals, with crown isola-
CCA suggests that sheet and orb web weavers were the only guilds tion limiting free exchange between trees (Southwood & Kennedy,
sensitive to island size (y‐axis), orb web weavers having their highest 1983; Adams et al., 2016). In this system, epiphytes behave like
diversity in smaller islands than sheet web weavers. Space and sheet islands within islands (Richardson, 1999). Our data confirm that
web weavers and ambush and other hunters were most sensitive to spider communities on epiphytes conform to the expected positive
island position (x‐axis), space web weavers having their highest relationship between island size and species numbers and that iso-
diversity in islands closer to the ground and the main trunk than the lation is also a weak but significant driver of spider diversity and
other three groups. Ground hunters were the guild closest to the similarity at the community level. The importance of canopy‐island
centre axis of the ordination plot suggesting a relatively low sensitiv- size for arthropod species numbers and abundance has been
ity for island size or isolation. reported previously (Borges & Hortal, 2009; Méndez‐Castro &
MÉNDEZ‐CASTRO ET AL.
-1.0
Log - island size
0.5
Space web weavers
CCA2
0.0
Ground hunters
F I G U R E 3 Canonical correspondence
analysis showing the ordination of spider
guilds in canopy islands (epiphytes) in a
-0.5
Mexican shade‐coffee plantation according
to the effect of the size and position of
the canopy islands. The effect of island
position is summarized by the interaction
Orb web weavers
between distance from the ground and
horizontal distance. R2 means proportion
-0.5
of variation explained
Rao, 2014; Stuntz et al., 2002). However, in contrast to our
results, the single other study addressing spatial distances among
epiphytes did not show significant effects of interisland distances
or isolation from the ground on epiphytic soil communities (War-
dle et al., 2003).
Within a tree canopy, microclimate, substrate characteristics,
and distance from the understorey vegetation, as a potential spe-
cies source, all differ with height above the ground. Consequently,
invertebrate diversity is generally found to change along the verti-
cal axis, but patterns are heterogeneous (Aikens & Buddle, 2012;
Gonçalves‐Souza et al., 2011; Pinzon et al., 2013; Rodgers & Kitch-
ing, 2011). For epiphyte‐dwelling spiders, the two available studies
related to vertical stratification report a decrease in species rich-
ness with height above the ground, tested up to 1.5 m (Gonçalves‐
Souza et al., 2011), and an increase with height above the ground,
tested up to 10 m (Méndez‐Castro & Rao, 2014). Our results,
including the full height of the trees, show that species richness
and the relative rareness per epiphyte island decreased with height
above the ground, although for species richness, this effect
depended on the horizontal position. This could be interpreted as a
negative effect of vertical isolation. However, with only about 5%
contribution of the spatial variables to the total variance in species
richness, compared to 68% explained by canopy‐island size, the
role of spatial gradients within the canopy appears to be smaller
than we expected. This is particularly surprising since trees tend to
have different epiphyte communities in different parts of the crown
(Wang et al., 2016; Woods, Cardelús, & De Walt, 2015), which
would, on top of differences in connectivity, add variation in the
living conditions for epiphyte‐dwelling spiders.
| 7
0.0 1.0
1.0
Sheet web weavers
Distance to ground
Horizontal distance
Ambush hunters
0.0
Other hunters
Island features R2
Island size 0.22
-1.0
Spaal posion 0.04
Residual 0.74
0.0 0.5 1.0
CCA1
Even though isolation had only a modest effect on spider species
numbers, distances between epiphytes clearly affected the species
composition. As expected by the ETIB, nearby islands were more
similar than islands at a larger distance from each other, up to a dis-
tance of about 4 m. Apart from providing shorter dispersal distances,
short distances between islands also imply a similar environment,
which may also partly explain species similarity. This is illustrated by
the increase in similarity at distances above 5 m, which is clearly not
attributable to easy dispersal between islands but most likely to the
similar microhabitat experienced at opposite positions in the outer
crown (Scheffers et al., 2014; Wang et al., 2016; Woods et al.,
2015).
The stronger effect of isolation (expressed as distance from
the ground or distance between islands) on the relative rareness
and on community similarity than on species numbers shows
that spatial features affect the species composition even if the
overall number of species remains similar. One likely reason for
the lack of overall isolation effects on diversity is indicated by
our ordination analysis, which points to spatial relationships
being specific for functional groups. Spider communities are very
heterogeneous mixtures of species differing in dispersal mode,
hunting behaviour, body size, and life‐cycle duration. According
to our results, analysing this whole mixture as one could mask
responses of guilds or other functional or taxonomic subgroups
to isolation and island size. At the small scale of epiphytes
within a tree, island size seems to have a more pronounced
effect on two out of the three web‐building spider groups, while
wandering errant hunters seem to be more affected by the ver-
tical and horizontal distribution of the islands. Even though an
8 |
exhaustive analysis of these lower level patterns is outside the
scope of this paper, this issue certainly deserves further atten-
tion (Mendez Castro et al., in prep) and should be taken into
account in future studies.
Epiphyte islands are relatively small and relatively close together
for most spider groups, compared to oceanic islands for most organ-
ism groups. Still, in both these systems, it is apparent that the basic
predictors of the ETIB (island size and isolation) are scale dependent
and taxon‐specific. For example, in the Aeolian islands, cursorial
invertebrates were unable to colonize neighbouring islands, while
birds were less sensitive to isolation (Fattorini, 2010). Such specific
effects of isolation on organisms with different dispersal abilities
vary across geographic scales and taxonomic groups, with examples
coming from different insular (Cardoso et al., 2009; Carvalho &
Cardoso, 2014; Fattorini, 2009; Meyer & Kalko, 2008), fragmented
(Moore, Robinson, Lovette, & Robinson, 2008; Öckinger et al.,
2009; Prugh et al., 2008), and microhabitat (Cobbold & MacMahon,
2012; Marini, Bommarco, Fontana, & Battisti, 2010) island systems.
The results of this study therefore not only support the idea of
epiphytic plants behaving like canopy islands but also provide
another example of how the relativity of isolation (experienced dif-
ferently depending on the mobility and behavioural traits of the
target organisms) is important across taxonomic groups and spatial
scales.
Three‐dimensional surveys of epiphytes are difficult to carry
out, but the information about the spatial organization of epi-
phytes provides valuable information needed for understanding
their ecology and for the design of sustainable harvesting methods
(Hietz, 2005; Toledo‐Aceves, Mehltreter, García‐Franco, Hernán-
dez‐Rojas, & Sosa, 2013). In this study, we analysed island size and
island isolation effects on species richness, although processes
leading to this pattern, immigration, and extinction could not be
addressed with the single sampling used. However, due to the
small spatial scale and relatively fast dynamics, epiphyte‐dwelling
biota present a very suitable system to study colonization and
extinction processes, which we recommend as a goal in further
research.
In summary, species richness on epiphyte islands increases with
island size conform the prediction of the equilibrium theory of island
biogeography, while isolation plays a minor role for species richness
but an important role in community composition. The relativity of
isolation due to different spatial behaviour of different spider taxa as
well as the processes leading to the observed patterns should be in
the focus of future studies.
ACKNOWLEDGEMENTS
We thank members of INBIOTECA for discussions, revisions, and
support, Verónica Hernández Arellano and Evaristo Calihua Chipa-
hua for their valuable help during fieldwork, and the anonymous
journal reviewers for helpful suggestions. We are grateful to the
brothers Martinez‐Gama (La Orduña) for permission to work on
their coffee plantations. This work was supported by an
MÉNDEZ‐CASTRO ET AL.
institutional fund to D. R. (Universidad Veracruzana, PROMEP/
103.5/10/7748), and a Ph.D. and Postdoc scholarship to F. E. M.
C. (CONACYT 307402/251146 &/251146). We are grateful to Dr.
Fernando Alvarez Padilla from Universidad Autonoma de Mexico
(UNAM) for the advice provided in the taxonomic identification of
the spiders.
ORCID
Francisco Emmanuel Méndez-Castro http://orcid.org/0000-0002-
8926-2490
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